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Herpetologists' League

A New Species of High Andean Anadia (Sauria: Teiidae) from Paramo el Riecito, Estado Trujillo, Venezuela Author(s): Enrique La Marca and Juan Elas Garca-Prez Source: Herpetologica, Vol. 46, No. 3 (Sep., 1990), pp. 275-282 Published by: Herpetologists' League Stable URL: http://www.jstor.org/stable/3892970 . Accessed: 15/04/2011 12:14
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Herpetologica,46(3), 1990, 275-282 ? 1990 by The Herpetologists' League, Inc.

A NEW SPECIES OF HIGH ANDEAN ANADIA (SAURIA:TEIIDAE) FROM PARAMO EL RIECITO, ESTADO TRUJILLO, VENEZUELA
ENRIQUE LA MARCA' AND JUAN ELIAS GARCIA-PtREZ2 lLaboratoriode Biogeografra,Escuela de Geograffa,Facultad de Ciencias Forestales, Universidad de Los Andes, Apartado Postal 116, Merida 5101-A, Venezuela 2Departamentode Biologfa, Facultad de Ciencias, Universidadde Los Andes, Merida 5101, Venezuela
ABSTRACT: A new species of lizard, Anadia hobarti, is described from the Paramo El Riecito, Estado Trujillo, in the Venezuelan Andes. The species belongs to the Anadia bitaeniata Group, an assemblage of high montane teiid lizards from the northern Andes. It is distinguished from other members of the group by the presence of interfemoral pores (located on ventral scales), its higher femoral pore count, the smaller number of scales between pore rows, the number of scales between adpressed limbs, and the presence of pale spots on its flanks. Key words: Reptilia; Sauria; Teiidae; Anadia; Anadia hobarti; Taxonomy; Paramo; Venezuela

ANADIA is a Neotropical genus of teiid lizards distributed from Costa Rica east to Venezuela and south to Ecuador. The genus is arranged currently into five species groups (Oftedal, 1974), three of which occur in Venezuela. Of the Venezuelan taxa, just two groups inhabit high montane environments: the A. marmorata Group, in the Venezuelan Cordillera de La Costa, and the A. bitaeniata Group, in the Cordillera de Meridaand the Venezuelan spur of the Cordillera Oriental de Colombia. The A. bitaeniata Group is represented in Venezuela by A. bitaeniata Boulenger, A. brevifrontalis (Boulenger), and A. pamplonensis Dunn. Two of these species (A. bitaeniata and A. brevifrontalis) inhabit some of the highest environments of the Andes of Estado Merida.Anadia bitaeniata occupies cloud forests at 2400 m in the Rio Mucujun valley up to 3100 m in the Paramo de La Culata (personal observation), whereas A. brevifrontalis inhabits paramos from 2900 m (Oftedal, 1974) to 3600 m (Lancini, 1968) in the Escorialand Mucubaji regions. In 1987, as part of a herpetological survey of Venezuelan p'aramosin the Cordillera de Merida, we collected 10 specimens of an undescribed species of Anadia from a paramo in the Estado Trujillo.Following the group characterization given by Oftedal (1974) and Harris and Ayala (1987), the lizards were assigned to the A. bitaeniata Group, in spite of the fact that 275

it has not been designated a unique derived charactersuggesting the monophyly of the assemblage. The object of this paper is to describe this taxon. Anadia hobarti sp. nov. Figs. 2-5 Holotype.-ULABG (Coleccion de Anfibios y Reptiles del Laboratorio de Biogeografia de la Universidad de Los Andes, Merida, Venezuela) 1715, an adult male from Paramo El Riecito, 2625 m, from near the headwatersof Rio Castan and Rio Riecito (a.k.a. Esdora), S of the village of Ortiz, about 15 km straight line SE of the city of Trujillo, Distrito and Estado Trujillo, Venezuela (about 9?14' N and 70?24' W: Fig. 1). Collected by E. La Marca, 18 April 1987. Paratopotypes (nine total).-ULABG 1713, with the same data as the holotype; ULABG 1719-1724, EBRG (Coleccion de Anfibiosy Reptiles de la EstacionBiologica de Rancho Grande, Maracay) 2131, and MCNC (Museo de Ciencias Naturales de Caracas) 6462, collected by Juan Elias Garcia Pe'rez and Enrique La Marca, 7 May 1987. Diagnosis.-The new taxon is distinguished from all other members of the bitaeniata group by the presence of 2-4 interfemoral pores located on ventral scales (Fig. 5) and, correspondingly,fewer scales (0-2) between the pore rows. The closest condition is that exhibited by Anadia pam-

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in A. hobarti, the interfemoral pores are ubicated on ventral scales. Furthermore, the femoral pore series in A. hobarti ocLAGO DE cupy almost the entire length of the thigh, MARACAIBO whereas in A. pamplonensis the pores are confined to the proximal portion of the thigh. Anadia hobarti has 11-13 total fem9 9 . ..... 100 50 100 oral pores (10-12 femoral and 1-2 interKmn femoral pores) at each side, a condition that is among the highest in the group, with the exception of A. bitaeniata which has from 7-11 femoral pores at each side. 8 8 Even when an overlap in number of pores occurs between these two species, the number of intermediate scales between the rows of pores are different. In A. hobarti, they range from 0-2, while in A. bitaeniata, they are from 6-8 (Table 1). 72 71 70 Adult females of Anadia hobarti have 7-9 scales between adpressed limbs, while FIG. 1.-Map showing localities in the Venezuelan Andes for species of the bitaeniata Group. Dashed adult females of A. bitaeniata have 2-3. line indicates border with Colombia. Symbols rep- Adult males of the new species have digits resent the species as follows: star, indicated by an that superimpose when limbs are adarrow, Anadia hobarti; square, A. brevifrontalis; cirpressed along body axis, while adult males cle, A. bitaeniata; triangle, A. pamplonensis. Dotted of A. brevifrontalis have five scales bearea indicates Venezuelan Andes above 2000 m. tween the adpressed limbs. Anadia hobarti has pale spots on flanks plonensis which has "inner femoral pores (Fig. 2), a character that has not been relocated preanally" (Harris and Ayala, ported elsewhere in the species group, ex1072 71

70

1987). The main difference between these two species is that those inner pores in A. pamplonensis are femoral in relationship (that is, ubicated on femoral scales), while

cept in A. pulchella. In A. pulchella, there


are 5-9 femoral pores, and there are no subocular scales protruding between supralabial scales, while in A. hobarti there

FIG. 2.-Anadia

hobarti holotype (ULABG 1715). Note pale spots on flanks. From a color slide by P. Soriano.

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FIG. 3.-Anadia

hobarti holotype ULABG 1715. Dorsal view of the head. Line

5 mm.

are 10-12 femoral plus 1-2 interfemoral poresand at least one subocularprotruding downward. A comparison of traits between A. hobartiand the remaining species of the A. bitaeniata Group is given in Table 1. Description of holotype.-Body slender; head enlarged posterolaterally (Fig. 3); nasal single, without groove below nostril; nasal not contacting rostral;nasal contacting loreal; frontonasal wider anteriorly, about same width as frontal, not contacting frontal; prefrontals one and a half times longer than wide; prefrontals contacting loreal, nasal, pre- and supraocular scales; eight supralabials; five enlarged infralabials; one sublabial; loreal contacts second and third supralabials on left side and second supralabial on right

side; prefrontalsmeeting to form a midline suture about one half length of prefrontals; one presupraocular; three supraoculars, the middle narrowestand smallest, contacting superciliaries; six superciliaries in unbroken row between supraoculars and ciliaries; superciliaries separated from ciliariesby one row of tiny scales;eight ciliaries of upper lid on left side and seven on right side; five large lower palpebrals, subhexagonal, slightly dusted with pigment; suboculars separated from palpebrals by two rows of tiny scales; three suboculars, subequal in size, two with an angular downward protrusion:one between fourth and fifth supralabials and anotherbetween fifth and sixth (Fig. 4); firstand third suboculars subequal in size and larger than other subocular(s);frontoparietalsslightly wider

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FIG.4- A

hb

th

o p

5 L e

ve

ot

ha

FIG. 4.-Anadia

hobarti holotype ULABG 1715. Lateral view of the head, left side. Line

5 mm.

posteriorly, one and a half times longer than wide, forming a long common suture; interparietal hexagonal, twice as long as wide, narrowerthan frontal,anteriorwidth slightly narrower than posterior edge; interparietal forming angular suture with occipitals and frontoparietals,anterior an-

gle being less than posterior angle; scales of first occipital row larger than those of second; scales of second occipital row larger than nape scales; ear opening with two rows of granularscales, the inner row with 10 and the outer row with five scales; horizontal length of ear opening about half

hobarti MCNC 6462. Ventral view showing femoral and interfemoral pores, and hemiFIG. 5.-Anadia penes. Pores stained with ink. Line = 5 mm.

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matrix of distinguishing traits among members of the Anadia hitaeniata Group. 1.-Comparative Abbreviations are: ALT = A. altaserrania, BIT = A. bitaeniata, BRE = A. brevifrontalis, HOB = A. hobarti, PAM = A. pamplonensis, PUL = A. pulchella.
Characters ALT BIT BRE HOB PAM PUL

Dorsal scales count Ventral scales count Midbody annulus scales count Scales between adpressed limbs (adults only)' Femoral pores Interfemoral pores Scales between femoral pores row Lateral spots Lip coloration Venter/palm-sole coloration Venter/subfemoral coloration Venter/subcaudal coloration Ventral scales

45-47 32-33 38 ? 3-4 0 ?16 absent spotted spotted/ ? pale spotted/ ?spotted spotted/ spotted IMBR.3

32-42 27-31 31-41 0 (&d) 2-3 (22) 7-11 0 6-8 absent dark dark/pale dark/pale dark/pale JUXT.4

40-50 29-37 30-38 5 (&3) 7 (99) 5-10 0 6 8 absent dark dark/dark dark/dark dark/dark JUXT.

42-48 31-35 36-40 0 (&8) 7-9 (22) 10-12 1-2 0-2 present pale dark/pale dark/pale dark/pale JUXT.

35-42 30-35 33-39 0 (&6) ? 5-8 0 4 absent pale pale/pale pale/pale pale/pale IMBR.

38-45 30-34 37-45 ? ? 5-9 0 ?8-14 present pale pale2/? pale2/pale pale2/pale IMBR.

I Scales were not counted when digits of the adpressed limbs were superimposed. The most common pattern is a pale background with two irregular rows of pale brown spots. IMBR. = Imbricated. JUXT. = Juxtaposed.

horizontallength of eye; single median chin shield followed by two pairs of large and one pair of small shields, anterior pair in broad medial contact, middle pair medially separated by two tiny scales; gular scales squarish;no gular crease or guttural fold; collar fold conspicuous; 10 enlarged scales in collar rows; lateral neck scales smaller than, but blending gradually with gulars and those of nape. Dorsal scales rectangular, smooth, unkeeled, one and a half times longer than wide, slightly rounded behind, arranged in 43 imbricate transverserows from posterior end of ear to base of tail; lateral scalessmallerand less regularin shape than dorsals;transverse rows of scales forming body annuli, except in the postaxillaryand preinguinal granular zones; 35 scales in midbody annulus; ventral scales rectangular, with rounded posterioredges, slightly larger than dorsals, arranged in 33 juxtaposedtransverserows from collar to vent; three transverse rows of rectangular enlarged scales constitutingthe preanalplate; eigth marginal preanal scales; 11 prominent femoral pores in left thigh and 12 in right thigh; two interfemoral pores; row of femoral and interfemoral pores separatedby two preanalscales;14 scales under fourth finger of left hand; 18 scales under

and 14 above fourthtoe of hind limb; scales of limbs pentagonal, rhomboid or subeircular, imbricate (except where granularon palms, soles, under digits, axillae and postfemoral surfaces); scales of tail rectangular, subimbricate anteriorly to juxtaposed posteriorly;dorsal tail scales smaller than ventral for full length of tail. Coloration in life (from E. La Marca Field Notes, April 1987).-Dorsum uniformly dark gray; flanks same as dorsum, with yellowish-cream spots; yellowishcream band along upper lip (Fig. 2); throat, venter, and ventral surface of tail yellowish-cream, with a slight green hue. Coloration in preservative.-Dorsum dark olive, without markings;flanks same as dorsum, with cream spots; gular region and lower part of supralabialsto tympanum, pale gray; ventral region from collar to cloacal opening highly pigmented with dark gray; ventral portion of tail cream; posterior part of thighs pale cream with scattered dark pigmentation; ventral surface of palms pale cream, with digits somewhat darker; dorsal surface of digits with alternate dark and pale scales. Measurements of holotype (in mm).Snout-vent length 86.1; tail length 151.2; head length 19.5; head width (at parietals) 13; comparative data between the holo-

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TABLE 2.-Comparative data of specimens of type series of Anadia hobarti (measurements in mm). SVL = snout-vent length, TL = tail length, HL = head length, HW = head width, FP = femoral pores, IFP = interfemoral pores, SBFPR = scales between femoral plus interfemoral pore rows, SBAL = scales between adpressed limbs (see Table 1). Sex SVL TL HL HW FP IFP SBFPR SBALd Museum No.

M M M M M M M M F F

86.8 86.1 82.4 78.3 70.0 67.2 59.4 54.2 83.0 68.0

151.2

99.5 95.8

20.6 19.5 18.2 17.3 15.1 14.1 13.0 12.0 16.4 13.0

15.0 13.0 13.8 12.2 11.0 10.2 8.5 7.2 10.0 8.0

11 11 10 11 11 10 11 10 11 11

(12)+ lc + IC (11).

(1O)a

1 1 1 2 1 2 1 1 1 (2)b 2

2 2 2 0 2 1 2 2 1 1

0 0 0 0 0 0 2 5 7 9

ULABG ULABG EBRG ULABG ULABG MCNC ULABG ULABG ULABG ULABG

1721 1715 2131 1720 1719 6462 1713 1723 1724 1722

Number in parentheses is femoral pore count on right side. Number in parentheses is interfemoral pore count on right side. Additional number indicates accessory pore: i.e., a femoral pore located below the main row of femoral pores and not considered as part of the latter. 11 Taken with forelimbs and hindlimbs adpressed to sides of body, with nails extended.

type and the paratypes are given in Table 2. Variation (n = 10).-Frontonasal narrower anteriorly in MCNC 6462, equal in width in ULABG 1720, contacting supralabial in EBRG 2131, ULABG 1719, 1723 (only on right side), and 1724; suture of prefrontal scales, one fourth to one half length of prefrontals;two supraocularsin EBRG 2131 and ULABG 1720; three to six superciliaries;one row of minute upper palpebrals in ULABG 1719 (only on right side) and EBRG 2131; ciliaries 6-8; 3-5 suboculars, protruding between (third, fourth), (fourth, fifth) and (fifth, sixth) supralabials;only one protrusioninto supralabials at each side of head in EBRG 2131 and ULABG 1723; 3-5 subhexagonal to quadrangular palpebral scales; only one row of tiny scales between palpebral and subocular scales in ULABG 1723; six infralabials (excepting five in right side of EBRG 2131 and ULABG 1722, five in left side of ULABG 1724, and seven in right side of ULABG 1723); none or 1-2 rows of ear opening granules, of variable number; horizontallength of ear opening from one-half to same length as the eye diameter; three to four pairs of chin shields; suture between second pair of chin shields absent to 3/4 times length of shield; interparietal shield subhexagonal to quadrangular, one and a half to two times narrower than frontal; collar row scales 8-13; 2-3 transverserows of enlarged scales posterior to collar fold.

Transverserows of dorsal scales, 42-48; 31-35 transverserows of ventral scales;3640 scales on midbody annulus; 6-8 scales line anteriorborder of vent; 1-2 interfemoral poresplus 10-12 femoral poreson each side; one accessory pore on each side in ULABG 1719 and 1720, located below the space between the first and second pore in the preanal region (Table 2); no preanal scale separating rows of interfemoral and femoral pores in ULABG 1720, but just one preanal scale in MCNC 6462 (Fig. 5), ULABG 1722, and 1724; 13-15 scales under fourth finger; 15-20 scales under, and 10-14 above, fourth toe. Sexual dimorphism.-Some secondary sexual charactersoccur in adult males but not in adult females, as indicated by the following proportionsand the corresponding t-test of the sample: (adult males n =
6, adult females n = 2, juvenile male n = 1); ratio head width/head length: adult

males (x = 0.72) versus adult females (x = 0.62), 0.001 < P < 0.01; adult females versus juvenile male (x = 0.60), P > 0.1; adult males versus adult females plus juvenile male (x = 0.61), P < 0.001. Scales between adpressed limbs: adult males (x = 0) versus adult females (x = 8), P < 0.001; adult females versus juvenile male (x = 5), P > 0.1; adult males versus adult females plus juvenile male (x = 7), P < 0.001. The subadult male (ULABG 1713) shows intermediate values between adult males and adult females for the preceding characters. Interfemoral and femoral

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pores: more conspicuous in adult males than in adult females and juveniles. Hemipenial morphology.-Hemipenis of paratype ULABG 1719 has 20 total flounces arranged symmetrically on each side; comb-like rows with subequal calcareous spinules; spinules on comb-like rows of proximal region opposite sulcus slightly larger and more separated than spinules on other rows; most proximal flounce lacking spinules and slightly smaller than the flounce above it; distal and middle flounces on each side chevronshaped,without a gap in spinule rowsalong downward-oriented apices; flounces increasing in width from tip to base of hemipenis; seventh to eleventh opposite-to-sulcus flounces interrupted in the midline apices; first to sixth opposite-to-sulcus flounces each forming an uninterrupted row of spinules; 16 most distal flounces bordering sulcus spermaticus,the remaining flounces next to absulcal midline region only; smooth area above the midline absulcal rows of contiguous spinules about one-fifth length of hemipenis; bulges of tip small, bilaterally symmetrical in the everted hemipenis. ULABG 1715 (holotype) has 19 flounces. ULABG 1721 has 15 flounces bordering sulcus and five flounces lying next to absulcal midline only; most proximalflounceapproximatelyone-fourth width of the one above it, with a short row of spinules at right side. ULABG 1723 has the most proximal flounce spinulated and less than one-fourth the length of the adjacent one. Natural history.-All specimens were collected under rocks in an open paramo ("Paramo Andino" of Monasterio, 1980). Rosette shrubs"frailejones"(Espeletia sp.) were the visually dominant plant species, with occasional congregations of tall caulirosulan Ruilopezia sp. The inversion of the vegetation belts in the type locality, resultingin a high Andean mountain forest above a lower open p'aramo vegetation, appears to be the result of colonization of deforested areas by pairamospecies. The largestsample (eight specimens) was taken on 9 May 1987, from 0830-1000 h, on a windy and misty day; temperatures fluctuated between 10 and 14 C. Three weeks earlier, on a sunny day with temperatures above 20 C, just two specimens could be

found between 1100 and 1300 h under rocks in the same place. The difference in the number of individuals under rocks on these two days seems to correspondto the activity periods I and III ("under rock retreats" and "moving outside rock retreats",respectively) reported by Swain et al. (1980) for Anadia brevifrontalis in the Paramo de Mucubaji, Estado Merida. A clutch of two eggs (ULABG 1714), each egg about 16 mm long by 10 mm wide, was found under a rock. A communal nest, such as those seen in another paramo species (e.g., Anadia brevifrontalis reported by Swain et al., 1980, and A. bitaeniata, personal observation) was found in one instance. This nest sheltered four pairsof eggs and two adult specimens of the new species. No other species of reptiles were found at the type locality. Several frogs (cf. Eleutherodactylus lanwere also taken at the cinii Donoso-Barros) type locality. One Eleutherodactylus shared a rock shelter with an Anadia hobarti. Anadia hobarti, like all Anadia lizards of the bitaeniata Group, has an evident sexual dimorphism in head size. Although we did not observeagonisticbehavior,there were indications suggestive of aggressive interactionsin the form of head scars and regenerated tails in about 75%of the sample of males and regenerated tail in one female (ULABG 1724) of two. The sexual dimorphism in head size, along with the head marksand regenerated tails, suggests that Anadia hobarti may have agonistic behavior. The first report of agonistic behavior in a species of the group was given by Fouquette (1968) for A. brevifrontalis. In A. hobarti, the hypothesized agonistic behavior may be the product of the scarcity of sheltering sites, which in paramo habitats may be a limiting resource. Etymology.-The specific name hobarti is a patronymic noun in the genitive singular,honoring Dr. Hobart MuirSmith, one of the world's greatest and most productive herpetologists.
Acknowledgments. -The senior author thanks Dr. H. M. Smith for his continuous counseling and warm friendship. Before dedicating the new species to him, Dr. Smith kindly read and commented upon an early version of this paper. P. Soriano and D. Cabello provided working space at the Grupo de Ecologia Ani-

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mal, Facultad de Ciencias, of the Universidad de Los Andes (ULA). P. Soriano graciously took the picture used in Fig. 2. The black and white picture for Fig. 5 was kindly taken by A. Sulbaran through the courtesy of Dr. E. Palacios Pru, of the Centro de Microscopia Electr6nica ULA. Dr. A. R. Lancini, Dr. H. M. Smith, and Dr. P. Vanzolini provided pertinent literature. Specimens of Anadia pamplonensis were examined through the courtesy of Dr. A. R. Lancini and Lic. P. Delgado (MCNC). A drawing showing femoral pores and femoral scales of the holotype of Anadia pamplonensis (ANSP 20835) was kindly provided by one anonymous referee. We also thank D. Borjas and M. Teran for camping assistance and companionship. The manuscript has greatly profited from the comments of the editors and three anonymous referees. All responsibilities remain ours, however. This work was supported in part by Grants S1-2053, from the Consejo Nacional de Investigaciones Cientificas y Tecnol6gicas de Venezuela (CONICIT), and Fo-223.89, from the Consejo de Desarrollo Cientifico, Humanistico y Tecnol6gico de la Universidad de Los Andes (CDCHT-ULA), to the senior author.

mos andinos. Ed. Univ. Los Andes, Merida, Venezuela. OFTEDAL, 0. T. 1974. A revision of the genus Anadia (Sauria, Teiidae). Arq. Zool. Mus. Zool. Univ. Sao Paulo 25:203-265. SWAIN, T. A., F. ARP, AND R. D. YOUNKIN. 1980. A preliminary report on the ecology of a tropical,

high altitude lizard, Anadia brevifrontalis.J. Herpetol. 14:321-326. Accepted: 6 December 1989

Associate Editor: John Iverson


APPENDIX I

Specimens Examined (34)


Anadia bitaeniata.-VENEZUELA: Estado Merida: Cloud forest above Monterrey, 2500 m, ULABG 1409; Paramo de La Culata, 3200 m, ULABG 16461647; cloud forest above Truchicultura de Monterrey, 2400 m, ULABG 1731; Asentamiento Campesino Monterrey, 7 km NE Merida, 2400 m, CVULA (Colecci6n de Vertebrados de la Universidad de Los Andes, Mrida) 4643-4645, MCNC 6514. Anadia brevifrontalis. -VENEZUELA: Estado Merida: Paramo de Mucabaji, CVULA-IV 471-477, CVULA-IV 765, 1568; Paramo de Mucubaji, 3400 m, CVULA-IV 4794; Morrena terminal, laguna de Mucubaji, Paramo de Mucubaji, ULABG 1535; Nr. La Venta, 3100 m, 17 km on road from Pico del Aguila to La Venta, ULABG 1569. Anadia marmorata.-VENEZUELA: Estado Aragua: Rancho Grande, ULABG 1514. Anadia pamplonensis. -VENEZUELA: Estado Tachira: Distrito Junin: Villa Paez, MCNC 6504, 6518. Anadia hobarti sp. nov.-VENEZUELA: Estado Trujillo: Paramo El Riecito, 2625 m, 15 km SE Trujillo, ULABG 1715 (holotype), and ULABG 1713, 1719-1724; MCNC 6462, EBRG 2131 (all paratopotypes); ULABG 1714 (clutch of two eggs).

LITERATURE
FOUQUETTE,

CITED

JR., M. J. 1968. Observations on the natural history of microteiid lizards from the Venezuelan Andes. Copeia 1968:881-884. 1987. A new HARRIS, D. M., AND S. C. AYALA. Anadia (Sauria: Teiidae) from Colombia and restoration of Anadia pamplonensis Dunn to species status. Herpetologica 43:182-190. LANCINI,A. R. 1968. El genero Euspondylus (Sauria: Teiidae) en Venezuela. Publ. Ocas. Mus. Cienc. Natur. Caracas, Venezuela; Zoologia 12:1-8.

MONASTERIO, M.

1980.

Las formaciones

vegetales

de los paramos de Venezuela. Pp. 93-158. In M. Monasterio (Ed.), Estudios ecol6gicos de los Para-